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Intertidal biota and washup at Ahipara and Herekino, Northland west coast.

Article · January 2004

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Bruce W Hayward Margaret S. Morley Geomarine Research Auckland Museum

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The user has requested enhancement of the downloaded file. All in-text references underlined in blue are added to the original document and are linked to publications on ResearchGate, letting you access and read them immediately. INTERTIDAL BIOTA AND WASHUP AT AHIPARA AND HEREKINO, NORTHLAND WEST COAST Bruce W. Hayward1, Margaret S. Morley2, Hugh R. Grenfell1, Rowan Carter3, Glenys C. Hayward4, Wilma M. Blom2, Doug Rogan5 1Geomarine Research, 49 Swainston Rd, St Johns, Auckland, 2Auckland War Memorial Museum, Private Bag 92018, Auckland, 3North Shore City Council, Auckland, 419 Debron Ave, Remuera, Auckland,5Canterbury Museum, Christchurch

SUMMARY

Three hundred and twenty-nine species are recorded from the Ahipara and Herekino Harbour shores of north-west Northland. These include 178 species of gastropods, 56 species of bivalves, 29 species of seaweeds, 14 species of crabs, 10 species each of chitons and barnacles, 9 species of echinoderms, 6 species each of anemones and polychaete worms. These records extend the recorded range of 12 mostly small gastropods. When the present observations are added to others from both coasts of northern New Zealand, we conclude that the boundary between the Cookian and Aupourian Marine Provinces would be better placed at the northern tip of Northland rather than near Ahipara.

INTRODUCTION

This study is one of several recently undertaken by the authors to document the poorly known diversity and biogeographic distribution of intertidal and shallow subtidal organisms along the west coast of the North Island of New Zealand. These are from north to south (Fig. 1): 1. Ahipara and Herekino Harbour (this study); 2. Whangape Harbour (Hayward et al., 1994); 3. Waimamaku Estuary (Hayward & Hollis, 1993); 4. Kawerua (Hayward, 1971, 1974, 1975, 1979, Hayward et al., 1995); 5. Waitakere Ranges (Hayward & Morley, in press); 6. Raglan Harbour area (Hayward et al., 2002); 7. Kawhia Harbour area (Morley et al., 1997); 8. Awakino to New Plymouth, north Taranaki (Hayward et al., 1999); 9. New Plymouth (Hayward & Morley, 2002).

Previous studies We know of no previous study that documents the intertidal life of the Ahipara or Herekino Harbour coast. Powell (1927) recorded 51 species of molluscs and described five new species from a fish trawl from 42 m depth (23 fathoms) off Ahipara, and Morley (1995) recorded the first west coast occurrence of Theora lubrica from Herekino Harbour.

Ahipara and Herekino Harbour Ahipara and Herekino Harbour (35º 10’ S) are situated on the west coast of northern Northland, west of Kaitaia (Fig. 1). Ahipara, at the south end of Ninety Mile Beach, has a broad sandy beach (Shipwreck Bay) with considerable shelter from the prevailing swells provided by Tauroa Pt. The intertidal rocky shore on either side of Mokurau Beach is also relatively sheltered, but exposure increases towards Tauroa (Reef) Point. The wide intertidal rocky platform along the west coast south of Tauroa Pt is fully exposed to the ferocity of Tasman Sea swells and storms. 15 km south of Tauroa Pt is the 200 m-wide entrance to the 6 km long, 0.4-1 km wide Herekino Harbour, clearly a drowned former stream valley. The shore on the north side of the harbour is largely sandy, but the southern coast near the entrance

Fig. 1. Location of intertidal and dredge study sites at Ahipara and Herekino, west Northland. consists of a relatively sheltered, rocky shore. The two branches of the upper harbour are largely composed of intertidal mud and sand flats surrounded by mangrove forest. At low tide the Hokianga Harbour is largely empty of water, with only a shallow subtidal entrance channel.

Field work Field work was undertaken by all the authors (except MM and DR) during three days of spring low tide in April 1999. The results of these studies have been supplemented by observations made by MM.

SPECIES LIST Mollusc nomenclature follows Spencer et al. (2002) and Marshall (2003).

Habitat where found: Qualitative assessment of abundance: A = Shipwreck & Mokurau Bch, washup a = abundant B = Shipwreck to Tauroa Pt, on rocks c = common C = south of Tauroa Pt, on rocks & washup f = frequent D = offshore, 40-45 m, dredged o = occasional E = Herekino Hbr, mangrove forest r = rare F = Herekino Hbr, Owhata, rocks & sand d = only seen dead G = Herekino Hbr entrance, rocks x = previous record H = Herekino Hbr, dredged subtidal * = extension of range A B C D E F G H A B C D E F G H CHITONS Eatonina subflavescens . . o . . . . . Callochiton crocinus . . d . . . . . Emarginula striatula . . d . . . . . Chiton glaucus . o f . . f f . Epitonium bucknilli . . . x . . . . Cryptoconchus porosus . r r . . . r . Epitonium jukesianum x d d . . . . . Eudoxochiton nobilis . r . . . . r . Epitonium minorum . . . x . . . . Ischnochiton maorianus . f f . . r o . Haliotis australis . . r . . . d . Leptochiton inquinatus . o ...... Haliotis iris d . d . . . d . Notoplax violacea . . r . . . . . Haliotis virginea . . o . . . . . Plaxiphora caelata . d r . . . . . Haminoea zelandiae . . d . . . . . Rhyssoplax stangeri . . r . . . . . Haustrum haustorium . o f . . . o . Sypharochiton pelliserpentis . c f . r f c . Herpetopoma bella . . d . . . . . GASTROPODS Herpetopoma larochei * . . d . . . . . Acteon cratericulatus . . . x . . . . Incisura lytteltonensis x ...... Aeneator attenuatus . . . x . . . . Janthina exigua . . d . . . . . Alcithoe fissurata . . . x . . . . Lamellaria ophione . . d . . . o . Amalda australis d ...... d Larochella alta * . . d . . . . . Amalda mucronata d ...... Lepsiella scobina . a f . . . f . Amalda novaezelandiae ...... Leuconopsis obsoleta . . d . . . . . Amphibola crenata . . . . a d . . Linopyrga rugata rugata . . d . . . . . Amphithalamus falsestea . o . . . . o . Lodderena formosa . . d . . . . . Anabathron hedleyi . c c . . . . . Lucerapex angustatus . . . x . . . . Antimelatoma ahiparana . . . x . . . . Macrozafra subabnormis . . d . . . . . Antisolarium egenum ...... d Maoricrypta costata * . . r . . . . . Aphelodoris luctuosa . . x . . . . . Maoricrypta monoxyla . o o . . . r . Aplysia dactylomela . o ...... Maoricolpus roseus manukauensis d r ...... Asteracmea suteri . d d . . . . . Marinula filholi . . d . . . . . Austrofusus glans d . d . . . d . Melagraphia aethiops . . d . c o . . Austrolittorina antipoda . a c . . f f . Merelina compacta * . . d . . . . . Austrolittorina cincta . c f . . o o . Merelina lyalliana . . d . . . . d Austromitra rubiginosa . o d . . . . . Merelina taupoensis * . . d . . . . . Hypermastus bulbula . . . x . . . . Mesoginella larochei . d d x . . . . Boreoscala zelebori d . . x . . . . Mesoginella pygmaea . d d . . . . . Bouchetriphora pallida . . d . . . . . Micrelenchus huttonii ...... d Brookula finlayi ...... Micrelenchus sanguineus . . d . . . o r Buccinulum linea . . d . . . . . Micrelenchus huttoni d ...... Buccinulum vittatum vittatum . d d . . . o . Microvoluta marginata . . . x . . . . Cabestana spengleri . o d . . . . . Neoguraleus amoenus . . d . . . . . Caecum digitulum . . d . . . . . Neoguraleus lyallensis ...... d Calliostoma punctulatum d . d . . . d . Neoguraleus murdochi . . d . . . . . Cantharidella tesselata . c c . . . c f Nerita atramentosa . o o . o o o . Cantharidus purpureus . . d . . . . d Notoacmea elongata . . f . . . . d Cellana ornata . o o . . . f . Notoacmea helmsi . . . . f f . d Cellana radians . c f . . f c . Notoacmea parviconoidea d o d . . . . o Cerithiella stiria . . d . . . . . Notoacmea pileopsis pileopsis . f d . . . . . Chemnitzia spp. . . d . . . . d Notoacmea helmsi f. scapha . . d . . . . . Cirsonella aff. laxa * ...... d Notoacmea scopulina ...... x . Cominella adspersa d . . . . o . . Notoacmea subtilis . . d . . . . . Cominella glandiformis . . . . o f d f Nozeba emarginata ...... d Cominella maculosa . . r . . . . . Odostomia takapunaensis . d d . . . . . Cominella quoyana quoyana . . d . . . . . Onchidella nigricans . . . . r . . . Cominella virgata virgata . d d . . . d . Onoba fumata . . d . . . . . Cookia sulcata . . o . . . . . Ophicardelus costellaris . . . . f . . . Crepidula youngi d ...... Paratrophon cheesemani . o ...... Crosseola vesca . . d . . . . . Paratrophon quoyi . . x . . . . . Cuvierina columnella . . . x . . . . Patelloida corticata . o o . . . o . Cylichna thetidis . . . x . . . . Penion sulcatus . . d . . . . . Cymatium parthenopeum . o . . . . o . Pervicacia tristis . d o . . . . . Dentomargo amoena . . . x . . . . Phenatoma rosea d ...... Dentomargo cairoma . . d . . . . . Pisinna rekohuana . d ...... Dicathais orbita . c o . . . f . Pisinna semiplicata * . . d . . . . . Diloma bicanaliculata . . r . . . . . Pisinna zosterophila . a d . . . d d Diloma coracina . . d . . . . . ?Pleurobranchea maculata ...... r . Diloma subrostrata . d d . c f . d Potamopyrgus estuarinus . . . . a . . . Diloma zelandica . . d . . . . . Proxiuber australe . . d . . . . d Doriopsis flabellifera . x x . . . . . Pupa kirki . . . x . x . . Eatoniella albocolumella . d o . . . . . Pusillina hamiltoni * . . o . . . . . Eatoniella flammulata . d d . . . . . Radiacmea inconspicua . d . . . . . d Eatoniella limbata . o ...... Ranella australasia ...... d . Eatoniella mortoni . . d . . . . . Retusa oruaensis x . d . . . . . Eatoniella notalabia . d ...... Risellopsis varia . d o . . . . . Eatoniella olivacea . c d . . . o . Rissoella cystophora . . o . . . . . Eatoniella pfefferi . c ...... Rissoella elongatospira * . . o . . . . . Eatoniella roseospira . . d . . . . . Rissoella rissoaformis * . . d . . . . . Eatonina crassicarinata? . . d . . . . . Rissoina chathamensis . d d . . . . . A B C D E F G H A B C D E F G H Rissoina zonata . . d . . . . . Nucula nitidula . . d . . . . . Sagenotriphora ampulla . . d . . . . . Panopea zelandica d ...... Scutus breviculus . o f . . . o . Paphies australis . . d . . c d a Seila cincta . . d . . . . . Paphies subtriangulata d . d . . . d . Semicassis pyrum . . d . . . d . Paphies ventricosa d ...... Serrata mustelina * . . d . . . . . Perna canaliculus . a o . . . o . Sigapatella novaezelandiae d . o . . . o . Peronaea gaimardi d ...... Siphonaria australis d d d . . . . . Philobrya munita . . d . . . . . Siphonaria propria d . d . . . o . Protothaca crassicosta d d d . . . d . Splendrillia otagoensis * . . . x . . . . Pseudoarcopagia disculus x . d . . . . . Stephopoma rosea . o d . . . . . Resania lanceolata d ...... ?Striodostomia orewa . d ...... Ruditapes largillierti x d ...... Struthiolaria papulosa d ...... Saccella bellula x ...... Syrnola crawfordi . . . x . . . . Scalpomactra scalpellum . d ...... Tanea zelandica d ...... Soletellina nitidula d ...... Taron dubius . o d . . . o . Spisula aequilatera d . d . . . d . Terelimella aupouria . . . x . . . . Talochlamys zelandiae . . d . . . r . Thoristella carmesina . . d . . . . . Tawera spissa d ...... Thoristella oppressa . . o . . . . . Theora lubrica . . . . . x . . Trimusculus conicus . . d . . . d . Trichomusculus barbatus . . d . . . . . Trivia merces . . d x . . . . Xenostrobus pulex d a a . r . a d Trochus viridis . . d . . . d . Zenatia acinaces d ...... Tugali elegans . . x . . . d . SCAPHOPODS Tugali suteri d . d . . . . . Antalis nana . . d x . . . . Turbo smaragdus . f f . r o . d CEPHALOPODS Uttleya ahiparana . . . x . . . . Argonauta argo . . d . . . . . Volvulella nesentus . . . x . . . . Spirula spirula d . d . . . . . Xymene traversi . d f . . . c . ECHINODERMS Zaclys sarissa . . d . . . . . Allostichaster polyplax . . r . . . . . Zalipais lissa ...... r Astropecten polyacanthus r ...... Zeacolpus ahiparanus d . . x . . . . Coscinasterias muricata . o o . . . . . Zeacumantus lutulentus . . . . . f . c Echinocardium caudatum ...... Zeacumantus subcarinatus . a f . . . f d Evechinus chloroticus . o o . . . r . Zebittium exile . . x . . . . . Fellaster zelandiae . . . . . d . o Zegalerus tenuis . d d . . . . d Ophionereis fasciata . r . . . . r . Zemitrella choava . . d . . . o . Patiriella regularis . o f . . . o . Zemitrella fallax . . d . . . . . Stichaster australis . o r . . . r . Zemitrella laevirostris? . . o . . . . . CRABS Zemitrella stephanophora . . d . . . . . Cyclograpsus lavauxi . . . . f f . . Zemitrella sp. . . d . . . . . Halicarcinus whitei . . . . . x . . Zeradina odhneri . . . x . . . . Helice crassa . . . . c . . . Zethalia zelandica d d . . . . . d Hemigrapsus crenulatus . . . . f . . . BIVALVES Hemigrapsus edwardsi . . r . . o o . Acar sandersonae . . x . . . . . Heterozius rotundifrons ...... r . Acar sociella . . d . . . . . Leptograpsus variegatus . f r . . . o . Anomia trigonopsis . r . . . . r . Macropthalmus hirtipes . . . . f . . . Arthritica bifurca ...... d Notomithrax ursus x ...... Austrovenus stutchburyi d ...... a Ovalipes catharus . . . . . d . . Bankia australis . . . . f . . . Ozius truncatus . f f . . . o . Barbatia novaezelandiae . . d . . . . . Pagurus novizelandiae . c f . . o f . Barnea similis d ...... Petrolisthes elongatus . f f . . o f . Bassina yatei d ...... Plagusia chabrus . f o . . . . . ?Benthocardiella obliquata . . d . . . . . BARNACLES Borniola reniformis . . d . . . . . Austrominius modestus . r . . c o . c Cardita aoteana . . d . . . . . Balanus variegatus . . d . . . . . Crassostrea gigas . o o . f f o . Calantica spinosa . r d . . . . . Divarilucina cumingii d ...... Chamaesipho brunnea . a f . . . c . Dosinia anus d ...... Chamaesipho columna . a a . . . a . Dosinia subrosea d ...... Epopella plicata . c f . . . o . Gari lineolata d ...... Lepas sp. . . d . . . . . Hiatella arctica . d d x . . . . Notomegabalanus decorus . d d . . . r . Irus reflexus . o d . . . r . Platylepas hexastylos x ...... Lasaea hinemoa . . d . . . . . Tetraclitella depressa . o o . . . . . Leptomya retiaria d ...... COELENTERATES Macomona liliana . . d . d . d c Actinothoe albocincta . o . . . . o . Mactra murchisoni d ...... Amphisbetia bispinosa d . d . . . . . Melliteryx parva . . d . . . . . Anthopleura aureoradiata ...... c Modiolarca impacta . r . . . . d . Diadumene neozelanica . . r . . . . . Modiolus areolatus . . d . . . d . Isactinia olivacea . c o . . r o . Moerella huttoni . . . x . . . . Isocradactis magna . f r . . . o . Montacuta semiradiata neozelanica . . d . . . . . Isoparactis ferax ...... r . Myadora striata d ...... POLYCHAETES Myllita stowei . . d . . . . . Flabelligera bicolor . o o . . . . . Myllitella vivens vivens d . x . . . . . Neosabellaria kaiparaensis . . f . . . . . Neolepton antipodum . d o . . . . . Pectinaria australis ...... d . Nucula hartvigiana . . d . . . . c Salmacina australis . o o . . . . . A B C D E F G H A B C D E F G H Spirorbis sp. . f f . . . . . Halopteris virgata . o o . . . . . Spirobranchus cariniferus . c a . . c c . Hormosira banksii . o f . . . o . PLATYHELMINTHES . r r . . . . . Landsburgia quercifolia . o o . . . . . SPONGES Laurencia thyrsifera . . o . . . . . Tethya aurantium . o ...... Lessonia variegata . o ...... ASCIDIANS ...... Melanthalia abscissa . o o . . . o . Corella eumyota ...... o . Osmundaria colensoi . f . . . . o . Pyura sp. . . o . . . o . Plocamium costatum . o . . . . o . SEAWEEDS Pterocladia capillacea . . o . . . o . Apophlaea sinclairii . . c . . . f . Pterocladia lucida . . o . . . o . Carpophyllum maschalocarpum . o o . . . o . ?Rivularia sp. . . o . . . . . Cheilosporum sagittatum . . o . . . . . Scytothamnus australis . o o . . . o . ?Cladophora sp. . o ...... Splachnidium rugosum ...... o . Codium convolutum . r r . . . o . Ulva lactuca . o ...... Codium dimorphum? . . o . . . . . Ulva sp...... o . Codium fragile ...... o . Xiphophora chondrophylla . o o . . . f . Colpomenia sinuosa ...... o . INTERTIDAL LICHENS Corallina officinalis . c c . . . c . Lichina confinis ...... r . Durvillaea antarctica d r r . . . . . INTERTIDAL VASCULAR PLANTS Ecklonia radiata . o o . . . o . Avicennia marina var. australasica . . . . a . . . Endarachne binghamiae . o . . . . o . Zostera muellerii . . r . . . . . Gigartina alveata ...... f .

HABITAT NOTES

Herekino Harbour The mid to high tidal mangrove forest that fills the upper reaches of both arms of Herekino Harbour supports the usual range of marine invertebrates. Most common invertebrates are the mud snail Amphibola crenata, topshells Melagraphia aethiops and Diloma subrostrata, and mud crab Helice crassa. Commonly attached to the trunks, lower branches and aerial roots are the acorn barnacle Austrominius modestus and rarer Pacific oyster Crassostrea gigas and flea mussel Xenostrobus pulex, while algal grazers in the branches include rare Nerita atramentosa, Turbo smaragdus, Onchidella nigricans, and Sypharochiton pelliserpentis. Among rushes around the upper fringes of the mangroves are common Potamopyrgus estuarinus and less frequent Ophicardellus costellaris. Thirteen invertebrate taxa were recorded living on and in the unvegetated fine to medium sand floor of Herekino Harbour and 20 more were present as dead shells (Appendix 1). Pipi Paphies australis, and cockle Austrovenus stutchburyi, are the dominant molluscs living both subtidally and intertidally (Fig. 2). The small sea anemone Anthopleura aureoradiata, are commonly attached to these two shallow-burrowing, infaunal bivalves, together with the acorn barnacle. Other common infaunal bivalves are the wedge shell Macomona liliana, nut shell Nucula hartvigiana, and occasional Theora lubrica. Common bivalves living on the sand surface are the horn shell Zeacumantus lutulentus, whelk Cominella glandiformis, and small topshell Cantharidella tesselata. Ninety-two rock-inhabiting taxa (81 living) were recorded from on and under rocks along the southern shore of Herekino Harbour between the road end at Owhata and the harbour entrance (Fig. 1). Zone forming organisms on the rocky shore are the periwinkle Austrolittorina antipoda, barnacles Chamaesipho brunnea and C. columna, the flea mussel Xenostrobus pulex, tube worm Spirobranchus cariniferus, and the algae Apophlaea sinclairii, Gigartina alveata, Xiphophora chondrophylla, and Corallina turf. Common grazers on the rocky shore include the limpets Cellana radiata and C. ornata, and the small seaweed- inhabiting Cantharidella tesselata, and carnivores include the white rock shell Dicathais orbita, oyster borer Lepsiella scobina, smaller Xymene traversi, and occasional specimens of the large Cymatium parthenopeum (Fig. 2). Fifteen additional species of seaweed were also recorded on the rocks, especially around the low tidal fringe. A rich fauna lives in the protection provided beneath low tidal cobbles, and includes numerous half crabs Petrolisthes elongatus, chitons Chiton glaucus and Ischnochiton maorianus, occasional gastropods Buccinulum vittatum, Taron dubius, soft-bodied Scutus

Fig. 2. Common intertidal and dredged species Herekino Harbour. Drawings by Powell (1987) and Margaret Morley. breviculus, sea squirt Corella eumyota, crabs Ozius truncatus and Hemigrapsus edwardsi, and brittlestar Ophionereis fasciata.

Open coast beaches Fifty-five species, mostly mollusc shells, were recorded washed up on the sand at the southern end of Ninety Mile Beach around Ahipara. Some of these are bivalves that live in the subtidal surf zone just offshore (e.g., Bassina yatei, Dosinia anus, Mactra murchisoni, Paphies subtriangulata, Spisula aequilatera); others have been washed in from deeper water during storms (e.g., Amalda mucronata, Antalis nana, Austrofusus glans, Gari lineolata, Myadora boltoni, Resania lanceolata, Soletellina nitidula, Tanea zelandica, Zeacolpus ahiparanus); some are pelagic (e.g., Spirula spirula); some are intertidal rock-inhabiting taxa (e.g., Calliostoma punctulatum, Haliotis iris, Tugali suteri); or have been carried by the tide out of

Fig. 3. Common or typical species of the open coast, west of Ahipara. Drawings by Powell (1979, 1987), Morton and Miller (1968) and Margaret Morley. a sheltered harbour and swept up the coast (e.g., Austrovenus stutchburyi, Cominella adspersa). During our visit, there was an unusually rich wash-up of 113 species on the high tidal beach above the low tidal reefs on the outside of Tauroa Pt. These included many of the rarer and unusual molluscs recorded in this study, most were derived from subtidal rock and sediment habitats.

Exposed rocky shore One-hundred and ten species were recorded live on or under the intertidal rocks from Ahipara round to Tauroa Pt. Grazing on the rocks above high tide are common periwinkles Austrolittorina antipoda and N. cincta, and on more shaded faces the limpet Notoacmea pileopsis (Fig. 3). Slightly lower on the shore are the zoning barnacles Chamaesipho brunnea, C. columna, and Epopella plicata, their predator Lepsiella scobina, together with the red leathery seaweed Apophlaea sinclairii, and grazers Cellana radiata and Sypharochiton pelliserpentis. In shallow, mid to high tidal pools are common horn shells Zeacumantus subcarinatus and olive anemones Isactinia olivacea. Forming patchy zones around mid tide are the flea mussel Xenostrobus pulex and tube worm Spirobranchus cariniferus. Towards low tide the rocks have a carpet of pink coralline turf and in places dense beds of the green- lipped mussel Perna canaliculus, with their predators Dicathais orbita, Haustrum haustorium, and the orange starfish Stichaster australis. Common micromolluscs in the seaweeds include Pisinna zosterophila, Cantharidella tesselata, Anabathron hedleyi, Eatoniella olivacea, and E. pfefferi. The most common crabs are the red rock crab Plagusia chabrus at low tide, the purple intertidal crab Leptograpsus variegatus, the black-finger crab Ozius truncatus, and half-crab Petrolisthes elongatus.

TYPES

Eight species of gastropod were described by Powell from off Ahipara. Seven are still considered valid and the eighth is now considered to be a junior synonym of an earlier described species. These species are: Off Ahipara 42 m Aeneator attenuata Powell, 1927 Antimelatoma ahiparana Powell, 1942 Balcis pervegrandis Powell, 1940 = junior synonym of Hypermastus bulbula (Murdoch and Suter, 1906) Finlayola crawfordi (Powell, 1927) as Syrnola crawfordi Uttleya ahiparana (Powell, 1927) as Rugobela ahiparana. Zeradina odhneri Powell, 1927 Zeacolpus ahiparanus (Powell, 1927) as Turritella (Zeacolpus) ahiparana Off Ahipara, 140 m Nodiscala ahiparana (Powell, 1930) as Pliciscala (Nodiscala) ahiparana

MOLLUSCAN BIOGEOGRAPHIC NOTES

The Ahipara and Herekino records of mollusc species extend the geographic ranges of 12 gastropods listed below (Fig. 4). Powell's (1979) published ranges have been used when commenting on extension of range, because Spencer and Willan (1996) give zoogeographic provinces only. These provinces (Powell 1979) are used to summarise the known ranges of each species (A = Aupourian, C = Cookian, F = Forsterian, M = Morioran, An = Antipodian). Additional records from the collections of the Auckland Museum (AK) and Margaret Morley (MM) are cited where they extend the published range.

* Cirsonella aff. laxa Powell, 1937, Skeneidae - previously recorded from depths of 260 m off the Three Kings, 90 m off the Snares Islands (Powell 1979), and 40 m off Pakiri, Northland (Morley et al. 1997b). The Herekino specimen (AK140828) is the first record from the west coast of the North Island. We have additional records from 90 m off Cape Maria van Diemen. These specimens show variation in the degree of separation of the body whorl and in details of ornamentation. There are 15 lots of Cirsonella in the Auckland Museum collections which are not identified to species level, they do not readily separate into the species described in Powell (1979). The Ahipara specimen is loosely coiled, the outer lip is strongly retractive to the suture and the umbilicus is about a third of the diameter of the base, all features of C. laxa, but it is smooth without the spiral chords and the body whorl is not completely separated from the aperture as described by Powell (1979). The range for Cirsonella laxa is now A and An provinces, including the east and west coasts of Northland.

* Herpetopoma larochei (Powell, 1926), Trochidae - previously recorded from the northeast of the North Island - Taupo Bay, Whangaroa, Great Barrier Island, Mayor Island (Powell 1979), Three Kings, and Whananaki, (AK), and at 1-15 m depth in the Bay of Islands (Morley and Hayward 1999). This Ahipara specimen washed up at Reef Point (AK 140599) is the first record from the west coast of the North Island.

* Larochella alta Powell, 1927, Aclididae - previously recorded from Mangonui, Awanui Heads, Rangaunu Bay, on the east coast of northern Northland, in depths of 10-20 m (Powell 1979). Also dredged from the Bay of Islands in 1-5 m (Morley and Hayward 1999). This specimen from Reef Point, Ahipara (AK 140601) is the first west coast record. The range is further extended by a specimen in anchor mud at 15 m depth in Dusky Sound, Fiordland (MM). The range for Larochella alta is now A, C and F provinces.

* Maoricrypta costata Sowerby, 1824, Calyptraeidae - previously recorded from Cape Maria van Diemen to Bay of Plenty (Powell 1979). Specimens from Reef Point, Ahipara (MM, AK) are the first west coast record. We also know of west coast North Island specimens from Waikawau and Kiritehere (MM) and the Waitakere coast (Hayward and Morley in press). The range is further extended by a specimen from Wharekauri, Chatham Islands (AK). The range for Maoricrypta costata is now updated to A, C and M provinces including the west coast of the North Island.

* Merelina compacta Powell, 1927, Rissoidae - previously recorded from off Mangonui, Northland in 10-20 m (Powell 1979), and known from Cape Maria van Diemen, Spirits Bay, and Rarawa, Northland; Poor Knights Islands in 40 m (MM) and Tryphena, Great Barrier Island (AK). These Ahipara specimens, washed up at Reef Point (AK140607), provide the first record from the west coast.

* Merelina taupoensis Powell, 1939, Rissoidae - this species is common in shell sand and under rocks at low tide on Northland's east coast from Cape Maria to Oneroa, Waiheke Island including Taupo Bay (Powell 1979), Colville (AK), and Great Barrier Island (MM). We have an additional record from algae wash at Sponge Bay, south of East Cape (MM). These Ahipara specimens (AK140610) provide the first west coast record. Thus the known range for Merelina taupoensis is now A and C provinces.

* Pisinna semiplicata (Powell, 1927), Anabathronidae - previously known from the northeast of the North Island, from Taupo Bay (Powell 1979) and the Bay of Islands (Morley and Hayward 1999). We have additional records from Spirits Bay; Great Barrier Island; Oneroa, Waiheke (MM); Tauranga; and Breaker Bay, Wellington (AK). The range for Pisinna

Fig. 4. Gastropods, whose recorded geographical range has been extended by this study. Drawings by Powell (1979, 1987) and Margaret Morley. semiplicata is now updated to A and C provinces, including the west coast with this record from Reef Point, Ahipara.

* Pusillina hamiltoni (Suter, 1898), Rissoidae - previously known from Tom Bowling Bay, Northland; Hauraki Gulf; Lyall Bay, Wellington; Banks Peninsula; Foveaux Strait; Waitangi, Chatham Island (Powell 1979); and the Bay of Islands (Morley and Hayward 2001). There are also specimens collected from East Cape (AK). These Ahipara specimens living on algae (AK140616), provide the first west coast record. This species is known from A, C, F, and M provinces.

* Rissoella elongatospira Ponder, 1966, Rissoellidae - previously known from the east coast of the North Island; Wellington west coast; east coast of the South Island; Stewart Island and Chatham Islands (Powell 1979). The range is already recorded from A, C, F, and M provinces (Spencer and Willan 1996) and further updated to include An province (Morley 1996). This specimen from Reef Point, Ahipara (AK 140647) is the first west coast record in Northland.

* Rissoella rissoaformis (Powell, 1939), Rissoellidae - previously recorded from the east coast of the North and South Islands, plus Fiordland and Stewart Island (Powell 1979), but was only recorded from C and F provinces by Spencer and Willan (1996). The range for Rissoella rissoaformis was previously updated with specimens from Auckland and Snares Islands (Morley 1996). The Ahipara specimens from algal wash (AK140620) are the first west coast record from the North Island. The provinces for Rissoella rissoaformis are now updated to A, C, F, and An.

* Serrata mustelina (Angas, 1871), Marginellidae - previously recorded from Northland east coast down to East Cape (Powell 1979). We know of specimens from off Raoul, Kermadec Islands in 10-30 m (Brook pers. comm.) and Mahia Peninsula (AK). The Ahipara specimen was collected at Reef Point. The range for Serrata mustelina is now A and C provinces, including the west coast of the North lsland.

* Splendrillia otagoensis Powell, 1942, Drillidae - previously known from off Oamaru, Otago in 30 m; and Paterson Inlet, Stewart Island in 20 m (Powell 1979). The Ahipara specimen (AK81489) was identified by F. Wells, a turrid expert. He has also identified S. otagoensis from Whangaroa in 29-31 m; Deep Water Cove, Bay of Islands, in 55 m; Whangarei Heads in 7 m; off Mangonui, Doubtless Bay in 146 m; and off Mayor Island in 154 m (AK). F. Wells has retained S. aoteana as a valid species by identifying specimens from off Cuvier Island and off Little Barrier Island (AK). The range for Splendrillia otagoensis is now updated to A and F provinces. This is the first west coast record in the North Island.

BIOGEOGRAPHIC PROVINCES

Six shallow marine biogeographical provinces have been established for the New Zealand region (Powell, 1955): Kermadecian (Kermadec Islands), Aupourian (north-east North Island), Cookian (central New Zealand), Forsterian (southern South, Stewart and Snares Islands), Moriorian (Chatham Islands), and Antipodean (Subantarctic Islands). There are several definitions of a biogeographical province. It may be defined as an area that exhibits a marked percentage of endemism; or the boundary between provinces coincides with the distributional boundaries of a significant number of taxa, or where there is a marked changed in the dominant taxa. Some modern workers reject the concept of provinces, as they are clearly artifical human inventions that oversimplify and hide the real complex nature of biogeographical distribution patterns. The relevance of biogeographical provinces to this Ahipara study lies in the fact that the western boundary between the Aupourian and Cookian Provinces was placed just north of Ahipara by Powell (1937), off Ahipara by Powell (1955), and between Ahipara and Kaipara by Spencer and Willan (1996). This boundary has presumably been placed off the west coast because of the presence of the weak southward-flowing, warm West Northland Current, which theoretically carries species southwards down the coast from the cape. Satellite imagery of water temperature suggests that the strength and distance it flows down the coast before meeting north-flowing currents is highly variable. The distance south that the province boundary has been placed has presumably been influenced by the recorded occurrences of some largely east coast molluscan species along the northern end of Northland’s west coast. Our study adds four, previously east coast-restricted mollusc species (Herpetopoma larochei, Merelina compacta, M. taupoensis, Serrata mustelina) to the small list that also occur as far south on the west coast as Ahipara. The total list with Ahipara as their southern limit is short, many have in recent years had their known ranges extended further south down the west coast by our studies, or even to the South Island. If we examine the list of 728 molluscan species recorded in all our recent intertidal and shallow subtidal studies on both the east and west coasts of northern New Zealand, we find that we have recorded 463 (64%) from both east and west coasts, with a further 200 only recorded from the east coast (27%) and 65 only from the west coast (9%). Going around the top of North Cape Peninsula is where by far the greatest change occurs in the molluscan fauna and it could be argued that this is a more logical place for the Province boundary. In our study data going southwards down the west coast, Ahipara is the southern limit for 40 species, Kawerua for 22 species, the Waitakeres for 52 species, Raglan for 20 species, and Kawhia for 18 species. Furthermore it could be argued that our intertidal and shallow subtidal records from Ahipara and Herekino have more in common with West Coast North Island (Cookian) than East Coast North Island (Aupourian) localities. The diversity of recorded molluscs from Ahipara (238 species) is only slightly higher than that recorded from Kawerua (192, Hayward et al., 1995), the exposed Waitakere coastline (194, Hayward and Morley, in press), Raglan (192, Hayward et al., 2002), Kawhia (188, Morley et al., 1997a) and north Taranaki coast (197, Hayward et al., 1999), and considerably less than that recorded from similar sections of coast in the Aupourian, such as Whananaki (360, in prep.), Waiheke (340, MM), and even Mahia (334, in prep.). This confirms the observations that a larger proportion of the warm- water Aupourian “endemics” are restricted to the east coast than those that extend down the west coast, and thus it could be argued that the boundary might indeed be more naturally placed at the northern tip of the North Island. Although many of the more common intertidal species are similar on both coasts, there are some clear differences, with the change occurring around North Cape area. For example, the bull kelp Durvillaea antarctica, the large orange starfish Stichaster australis, the colourful camouflaged anemone Isocradactis magna, and the gastropod Paratrophon cheesemani, are characteristic elements of all the exposed shores along the west coast (including Ahipara), but are all but absent from the East Coast. Dense intertidal beds of the green-lipped mussel Perna canaliculus are also a dominant zoning element on west coast shores, but they are far less abundant on the east. Two of the more common zoning organisms on east coast shores are the snail Nerita atramentosa and the brown seaweed Xiphophora chondrophylla, both of which are rare on the west coast with their highest west coast abundances in the north around Ahipara. Two other common molluscs on the east coast that are all but absent on the west are the slipper limpet Maoricrypta costata and Venericardia purpurata. As the above discussion confirms, the province boundary is somewhat artificial and drawing it as a thin line hides the fact that the faunal changes are more gradual and spread over a broad zone, reflecting the complex and fluctuating physical oceanographic patterns and the differing tolerance limits and niche requirements of every species.

ACKNOWLEDGMENTS We thank Jessica Hayward for drafting and F. Wells for identifying the Splendrillia species.

REFERENCES Hayward, B.W. 1971: Some factors affecting zonation of rocky shore organisms at Kawerua. Tane 17: 137-148. Hayward, B.W. 1974: Kawerua crabs. Tane 20: 158-161. Hayward, B.W. 1975: Kawerua echinoderms. Tane 21: 59-60. Hayward, B.W. 1979: An intertidal Zostera pool community at Kawerua, Northland and its foraminiferal microfauna. Tane 25: 173-186. Hayward, B.W.; Hollis, C.J. 1993: Ecology of Waimamaku River estuary, north of Kawerua, North Auckland. Tane 34: 69-78. Hayward, B.W.; Morley, M.S. 2002: Intertidal biota of the proposed Nga Motu Marine Reserve, New Plymouth. Poirieria 28: 1-11. Hayward, B.W.; Morley, M.S. in press. Intertidal life around the coast of the Waitakere Ranges. Auckland Regional Council Technical Publication. Hayward, B.W.; Blom, W.; Morley, M.S.; Stephenson, A.B.; Hollis, C.J. 1994: Benthic ecology of Whangape Harbour, Northland. Records of Auckland Institute and Museum 31: 219- 230. Hayward, B.W.; Morley, M.S.; Riley, J.; Smith, N.; Stace, G. 1995: Additions to the mollusca from Kawerua, North Auckland. Tane 35: 183-193. Hayward, B.W.; Morley, M.S.; Stephenson, A.B.; Blom, W.; Grenfell, H.R.; Prasad, R. 1999: Marine biota of the north Taranaki coast, New Zealand. Tane 37: 171-199. Hayward, B.W.; Morley, M.S.; Blom, W.; Grenfell, H.R.; Smith, N.; Rogan, D.; Stephenson, A.B. 2002: Marine biota of Raglan, Waikato West Coast. Poirieria 28: 1-16. Marshall, B.A. 2003: A review of the Recent and late Cenozoic Calyptraeidae of New Zealand (Mollusca: Gastropoda). The Veliger 46: 117-144. Morley, M.S. 1995. Extension of range for Theora lubrica. Poirieria 17: 21-22. Morley, M.S. 1996: Marine Molluscs of the Auckland and Snares Islands, Southern New Zealand January 1995. Poirieria 18: 2-25. Morley, M.S.; Hayward, B.W.; Stephenson, A.B.; Smith, N.; Riley, J. 1997a: Molluscs, crustacea and echinoderms from Kawhia, west coast, North Island. Tane 36: 157-180. Morley, M.S.; Goulstone, J.; Hazelwood, B.; Poortman, P.; Smith, N.; Stace G.; Thompson, F. 1997b: The last word on Mission Bay. Poirieria 21: 18-26. Morley, M. S.; Hayward, B. W. 1999: Inner shelf mollusca of the Bay of Islands, New Zealand, and their depth distribution. Records of the Auckland Museum 36: 119-138. Morton, J.E.; Miller, M.C. 1968: The New Zealand Sea Shore Collins, 653pp. Powell, A.W.B. 1927: Mollusca from twenty three fathoms off Ahipara, New Zealand. Transactions and Proceedings of the New Zealand Institute 58: 295-300. Powell, A.W.B. 1937: New species of marine Mollusca from New Zealand. Discovery Reports 15: 153-222. Powell, A.W.B. 1940: The Marine Mollusca of the Aupourian Province New Zealand. Transactions of the Royal Society New Zealand 70: 205-248. Powell, A.W.B. 1955: New Zealand biotic provinces. Conchology Section of the Auckland Museum Bulletin 11: 10-14. Powell, A.W.B. 1979: New Zealand Mollusca. Collins, 500 pp. Powell, A.W.B. 1987: Native Animals of New Zealand. Auckland Institute and Museum, 88 pp. Spencer, H.G.; Willan, R.C. 1996: The marine fauna of New Zealand. Index to the fauna: 3. Mollusca. New Zealand Oceanographic Institute Memoir 105. Spencer, H.G.; Willan, R.C.; Marshall, B.A.; Murray, T.T. 2002: Checklist of the Recent Mollusca described from the New Zealand Exclusive Economic Zone. http://toroa.otago.ac.nz/pubs/spencer/Molluscs/index.html

Appendix I. Census data for 10 litre dredge samples taken inside Herekino Harbour Dredge penetration averaged 0.1 m into the sea floor sediment. Live organisms are in numbers of individuals; presence of dead shells is indicated by d. See Fig. 1 for locations.

L261.. 70 71 72 73 74 75 L261.. 70 71 72 73 74 75 GASTROPODS ...... Turbo smaragdus . . . . d . Amalda australis . . . d . . Zalipais lissa . . . . . 1 Antisolarium egenum . . . d . . Zeacumantus lutulentus 4 d 1 d 3 d Cantharidella tesselata . . . . 11 . Zeacumantus subcarinatus . . . . d d Cantharidus purpureus . . . . d . Zegalerus tenuis . . . d . . Chemnitzia spp. . . . d d d Zethalia zelandica . . . . d d Cirsonella aff. laxa . . . . d . BIVALVES ...... Cominella glandiformis . d 5 1 2 d Arthritica bifurca . . . d d . Diloma subrostrata . . . d d d Austrovenus stutchburyi 193 1 . d 12 d Merelina lyalliana . . . . d . Macomona liliana 1 . 4 d d 1 Micrelenchus huttonii . . . . d d Nucula hartvigiana 3 . 1 d 1 . Micrelenchus sanguineus sanguineus . . . . . 1 Paphies australis 1 1.5 39 19 3 11 Neoguraleus lyallensis . . . . d . Xenostrobus pulex . . . d d d Notoacmea elongata . . . d d . ECHINODERMS ...... Notoacmea helmsi d d . . . d Fellaster zelandiae . . . . 2 d Notoacmea parviconoidea . . . d 1 d BARNACLES ...... Nozeba emarginata . . . d . . Austrominius modestus . . . 6 3 . Pisinna zosterophila . . . . d d COELENTERATES ...... Proxiuber australe . . . . d . Anthopleura aureoradiata . 5 2 4 38 . Radiacmea inconspicua . . . . d .

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